Spring 2012 A Publication of the Native Society Volume 3, Issue 1

Art by Tina ieme Brown White Oak (Quercus alba L.) Beech Family (Fagaceae) Official State Tree of Maryland

Tina ieme Brown Letter from the President A Publication of the Dear Members, Maryland Native Plant Society For the first issue of 2012, I’ll give you an overview of the Society’s activities in 2011. We started what we hope will be a tradition, to focus on a single plant group for the entire year. We picked ferns, those ancient, ubiquitous, and hard-to-distinguish . Ferns were featured in seven field trips, three monthly meetings, and two issues of Marilandica. Dwight Johnson’s Key to the 25 Common Ferns of Maryland proved indispensable throughout the year; it’s available on our website. Members responded enthusiastically, and for 2012 we’re focusing on Oaks. www.mdflora.org Field trips are our core activity. Last year the Society sponsored over 30 field trips, plus a P.O. Box 4877 Silver Spring, MD 20914 field excursion to the Pocono Plateau, and additional field trips at the conference. We thus enabled hundreds of people to experience the deep satisfaction of going outside and paying close CONTACTS attention to what we see. anks to Cris Fleming and Bob Yacovissi, plant lists from many field Membership trips are now posted on the website. Attendance at our meetings and field trips is at an all-time Karyn Molines, [email protected] high. e annual fall conference is a highlight of the year. Our 2011 Conference at LaPlata in Marilandica Editor Charles County focused on Southern Maryland’s rich plant communities and the urgent necessity Carolyn Fulton, [email protected] to protect them from encroaching development. Watch for announcements of this year’s Confer- Webmaster ence to be held at Towson University under the leadership of the Greater Baltimore Chapter. Iris Mars, [email protected] We continue to advocate for conservation. Carol Jelich testified in favor of legislation General Inquiries [email protected] (enacted) that will regulate the sale of invasive non-native plant species. Linda Keenan testified at a hearing on Prince Georges County’s Preservation Area Functional Master Plan. Kirsten MNPS CHAPTERS Johnson and Mary Pat Rowan were invited to speak at a US National Arboretum stakeholders Catoctin meeting. Linda Davis, Kirsten Johnson, Eugene Meyer, Bob Stanhope and other members Jim and Teresa Gallion, [email protected] participated in the Coalition for Responsible Deer Management, described in the Conservation Joe Metzger, [email protected] Watch section. In addition, we often cosign petitions and testimony by other organizations on Eastern Shore Leslie Hunter-Cario, [email protected] conservation issues. Greater Baltimore e Greater Baltimore Chapter welcomed Chris Partain as its new Chair. Outgoing Chair Christa Partain, [email protected] Ann Lundy, a founding member of the chapter, will continue on the chapter’s board. e Montgomery County chapter is now holding meetings monthly in Baltimore City, Baltimore County, and Howard [email protected] County. Under Liz McDowell’s leadership, our Western Mountains Chapter continued its regular North East meetings with speakers and outreach activities. In Montgomery County, some of our monthly Matthew Bazar, [email protected] meetings have been standing-room-only, so please arrive early to get a seat and to socialize. Prince George’s/Anne Arundel Counties For details of all our activities in 2011, please see our 2011 Annual Report in the “About” Matt T. Salo, [email protected] section of our website. Southern Maryland Karyn Molines, [email protected] ~ Kirsten Johnson Washington, DC Mary Pat Rowan, [email protected] Western Mountains Liz McDowell, [email protected] Western Mountains Chapter EXECUTIVE OFFICERS Kirsten Johnson, President e Western Mountains Chapter is celebrating the Year of the Oak with assistance from Wade [email protected] Dorsey, Savage River State Forest manager. Wade will lead a leisurely walk in June to highlight Marney Bruce, Vice President Linda Keenan, Vice President various oak species found in Garrett County. en as an encore at our October meeting, he will Marc Imlay, Vice President share his presentation on Maryland’s native oaks. Ginny Yacovissi, Secretary Our chapter meetings this year cover a range of topics. At our February meeting Liz Matt Cohen, Treasurer McDowell, Chapter Coordinator, explained how gardeners can play a key role in maintaining regional biodiversity by ‘going native'. In April we will focus on pawpaw, as plant breeder, R. BOARD OF DIRECTORS Neal Peterson, recounts its history, biology, and his recent efforts at its domestication. At our Ken Bawer Karyn Molines Matthew Bazar Christopher Puttock June meeting the guest speaker will be Donna Ford-Wentz, Herbarium Curator, West Carole Bergmann Glenn Rice University. Her main research interests include the flora of and plant collections Melanie Choukas-Bradley Mary Pat Rowan management. In August Stephen Keller, Assistant Professor, Appalachian Laboratory, will Cris Fleming Roderick Simmons discuss his current research on the ecology and evolution of invasive species and the conservation Carolyn Fulton Gary Steele genetics of rare and declining species. Albert Hartley Lou Aronica, Emeritus Our chapter members continue to spread the word about the value of native by serving Brett McMillan Joe Metzger, Emeritus on Frostburg University’s “Tree Campus USA” Advisory Committee and Arboretum Task Force, assisting local garden clubs with questions and representing MNPS at local native plant sales. ~ Liz Mc Dowell, Western Mountains Chapter Coordinator.

Marilandica Spring 2012 page 1 Oak in Focus—White Oak

Quercus alba L. gray, glabrous or nearly so when mature (greenish or reddish and may Beech Family (Fagaceae) be hairy when young), with scaly, reddish brown, ovoid or nearly by Melanie Choukas-Bradley round winter buds. Growth Habit: Large tree with a full, rounded crown. Woodland uring the “Year of the Oak,” the Maryland Native Plant trees are tall and straight; trees grown in the open may have a short, Society is celebrating the 21 native oaks of Maryland and wide trunk and broadly spreading limbs. we are learning all we can about their identifying charac- teristics and their importance to our regional ecology. Similar Species: Although Quercus alba is by far the most common of Surely none of our 21 species is more beloved than the the trees in the white oak group growing in Maryland, it could be white oak (Quercus alba), our official state tree. Abun- mistaken with other species. Chestnut oak (Q. prinus) has more dant in woodlands throughout Maryland, the white oak has shallowly lobed leaves and bark that is dark, thick, and separated into o long been a friend to humankind, both as a forest and ridges and valleys. Swamp chestnut or basket oak (Q. michauxii) has cultivated tree. Montgomery County Forest Ecologist and leaves similar to chestnut oak, and bark similar to white oak. It is MNPS board member Carole Bergmann speaks for all of us when she largely a tree of the coastal plain although a few specimens grow at says: “Such a beautiful tree! e pale gray shingled bark stands out so Blockhouse Point and in other piedmont locations and it is planted in beautifully against a bright blue winter afternoon sky. e white oak the Washington, D.C. area. Swamp white oak (Q. bicolor) has thick is our state tree for a reason. I love that tree and always point it out on leaves with felt-like pubescence below and long-stalked acorns. Post every walk I lead!” White oak acorns are edible after treating to oak (Q. stellata) has leaves with a large pair of lobes near the apex, remove tannin. Traditionally, a protein-rich meal was made from the giving the leaf a cross-shaped look. See www.mdflora.org for a full list of crushed, ground acorns and used in baked goods. e acorns are an oaks native to MD and other resources related to the Year of the Oak. important food source for wildlife, including squirrels, raccoons, Locations: e white oak can be found growing in woodlands white-tailed deer, wild turkeys, and quail. Less well-known is the throughout Maryland and it is a favorite cultivated tree. According to importance of oaks to , including butterflies and moths. Botanist, Ecologist and MNPS board member Rod Simmons: “Most MNPS President Kirsten Johnson observes: “Doug Tallamy says oaks of the mature White Oaks we see in yards and along old neighbor- support 534 Lepidopteran species – more than any other in the hoods and sections of cities are remnants of native forested areas midatlantic. us oaks also support -eating birds and .” dominated by upland oaks that were present when the houses and e white oak is, not surprisingly, in the white oak group, along with streets were built.” eight other Maryland oak species. Trees in the white oak group bear leaves with rounded (non-bristle-tipped) lobes or teeth (or apices in reats to the White Oak and other Oaks: Kirsten Johnson notes non-toothed or lobed species) and acorns maturing in one year. e that the white oak and other oaks are threatened by deer and urban- acorns in the white oak group are purportedly “sweeter” than those in ization. On the latter, Rod Simmons agrees: “[ e white oak] can the red oak group. tolerate the successive periods of drought and heat associated with climate change, but it cannot tolerate significant water table drops Native Habitat and Range: A variety of habitats including upland and loss of groundwater infiltration around its root zone that result woods; grows best in deep, rich, well-drained soils. Eastern U.S. and from the now-constant practices of digging, trenching, paving, over- southeastern from Maine and southern Quebec to Minne- building, burying underground utilities, etc., that are the hallmarks sota, south to eastern and northern . of our over-developed and highly fragmented suburbs and neighbor- Leaves: Simple, alternate, deciduous. 3 ½ to 9 inches (9-22.8 cm) hoods.” Geologist and tree enthusiast Tony Fleming adds: “I think long. Some leaves have narrow lobes separated by sinuses cut almost Rod hit the nail on the head: local environmental change is a to the midrib; others have wider lobes and sinuses cut only about well-documented culprit in plant and mortality, and it comes halfway to the midrib. Five to nine rounded lobes; lobes point in a variety of packages, large and small. To my thinking, the most upward and sometimes have one or more large, rounded teeth. Base pervasive form of local environmental change is deforestation and wedge-shaped or slightly rounded. Pubescent at first, soon becoming urbanization: the [D.C.] metro area probably has considerably fewer glabrous; very pale and sometimes slightly glaucous below. Petiole ¼ forested areas now than 50 or 100 years ago. Regional deforestation to 1 inch (0.5-2.5 cm) long. Autumn color: deep wine-red some creates a hotter, drier, and more wind-prone microclimate, not to years. Dry leaves often remain on the tree through the winter. mention less overall soil moisture availability as forested areas are converted to urban land cover. For the large trees that remain, the Flowers: Male flowers in loose catkins sometimes called “aments.” cumulative effect of all these local environmental changes amounts to Female flowers are tiny, with three-lobed stigmas. Trees are monoe- death by 1,000 cuts.” cious and spring-blooming. Fruit: Acorn, maturing during the first year as all acorns in the white Oak in Focus is adapted from City of Trees: e Complete Field Guide oak group do. Sessile or short-stalked. ½ to 1 inch (1.3-2.5 cm) long, to the Trees of Washington, D.C., Melanie Choukas-Bradley with enclosed for about one-quarter of its length by a bowl-shaped cup illustrations by Polly Alexander (University of Virginia Press). Carole covered with thickened nubbly scales. Bergmann, Charlie Davis, Cris Fleming, Tony Fleming, Kirsten Bark and Twigs: Bark pale ash-gray, with shaggy vertical scales; often Johnson, Wesley M. Knapp, Rod Simmons, and Tina ieme Brown slightly furrowed toward the base of the tree. Twigs reddish brown or contributed to this article.

Marilandica Spring 2012 page 2 On January 13, the Baltimore Sun reported that Governor O’Malley is visible, threat to our parks and other natural areas, namely the destruction of including in his proposed capital budget almost $24 million for much needed habitat due to the overpopulation of white-tailed deer. We sent the following renovations and improvements to infrastructure in the state parks. MNPS letter to Governor O’Malley, and we were pleased to receive this thoughtful supports this investment. e deterioration of buildings, piers, and parking response (facing page) from John R. Griffin, Secretary of the Department of lots is visible to everyone. But we wanted to call attention to another, less Natural Resources.

Sent through the Governor’s website: January 21, 2012

Dear Governor O’Malley, Maryland Native Plant Society supports your recently announced capital budget proposal to increase funds for needed improvements and maintenance in our state parks. But we want to call your attention to another, even more effective and likely less costly, way to improve Maryland’s parks. Nothing is causing more harm to our state parks, forests and natural areas than the exploding population of white-tailed deer. In many parks, the native shrub and herbaceous plant layer has been virtually eliminated. When the large trees in many of our forests fall and die, they are not being replaced by younger trees because white-tailed deer eat the tree seedlings and saplings before they have a chance to mature. You can walk miles in our woods and not see a single oak, hickory or maple sapling. Our forests are dying a slow death. e overpopulation of white-tailed deer encourages the proliferation of non-native invasive plants that the deer do not eat. ese non-native plants fail to provide food for the animals, birds, amphibians, and insects that depend on native plants for food and shelter. us, the entire forest ecosystem is harmed. In addition, deer threaten public health and safety by carrying Lyme disease and causing thousands of vehicle collisions annually in Maryland alone. Studies have shown that the white-tailed deer population must be reduced to fewer than 20 deer per square mile for forests to regen- erate. Yet the deer population in some areas of Maryland, such as Baltimore County, is estimated at more than 80 deer per square mile. Despite the destruction that white-tailed deer are causing, they are still protected by a regulatory system established decades ago when the deer population was sparse. State law and policy should now be updated. Park managers should be permitted — and encouraged through adequate funding — to control deer populations through managed hunting. In addition, white-tailed deer could, like the fish and crustaceans of the Chesapeake Bay, be managed as an economic resource. To this end we propose that the commercial sale of wild-hunted venison be permitted under an appropriate regulatory system that addresses public safety and balances the interests of recreational and commercial hunters. We believe this would be a practical way to reduce the deer population to a healthy level while at the same time providing revenue for our licensed hunters, and healthy high-protein, low-fat food for the people of Maryland. ank you for your consideration of our view. Yours truly, Kirsten Johnson, President

Early Spring at Robert E Lee Memorial Park

Buds of Quercus marilandica, anonyma Rosette of Arabis lyrata, Blackjack Oak (syn: Senecio smallii) Lyre-leaved Rockcress

Rosette and bud of Leaves of Quercus stellata, Saxifraga virginiensis, Post Oak Early Saxifrage

Marilandica Spring 2012 page 3 Ms. Kirsten Johnson 550 W. University Parkway Baltimore MD 21210

Dear Ms. Johnson:

Thank you for your email to Governor Martin O’Malley regarding deer management in Maryland’s State Parks. Governor O’Malley has reviewed your email and asked me to respond on his behalf. I appreciate you taking the time to express your support for the proposed increase in capital funding for State parks. With the support of the General Assembly in approving the Governor’s proposal, we will be able to address some long-standing capital improvement needs and look forward to the enhancements they will bring. I was interested to read your description of the damage deer are doing to many of our public lands. We have been well aware of the impacts caused by too many deer and have been working very hard to address this issue for over two decades. As you may remember, some of our early efforts to control deer on State parks resulted in significant protests from the general public. We met this resistance with a goal of education and compromise and have since moved to an era where the necessity for deer control is understood, and accepted, by most citizens. As a result, we now have managed hunting for deer in most of the parks where this activity is necessary and appropriate. While there may be the need for additional hunting on some tracts, we certainly feel our management philosophies have evolved to be consistent with the growing deer population. Consequently, the deer hunting opportunities on our State parks are very liberal, as we agree that managed hunting is the best means to manage white-tails. Gone are the days when hunting in a State park was considered an anomaly. Looking at deer harvests across our State, it is evident that hunters are doing a good job as deer managers. That harvest now includes far more antlerless deer (does) than antlered deer (bucks), a significant change from just a decade ago. Taking more does is the most effective way to reduce the herd, and as a result, we have seen the statewide deer herd drop since peaking in 2002. Our deer managers continue to make changes to liberalize hunting regulations to increase harvest and the good news is we have turned the corner and our deer herd is no longer growing. Your suggestion for establishing a commercial outlet for wild venison is not a new one, but it certainly is an uncommon one. As you may know, more and more people are looking at wild venison as being a healthy, sustainable and local source of food. I can point to a number of people that have taken up the sport of deer hunting, not because it was a family tradition, but because it provides an excel- lent source of food with an ecologically desirable outcome – fewer deer. The Department of Natural Resources has several programs in place to keep this trend growing; we look forward to adding to our deer hunting community every chance we get. I will give your suggestion some real consideration and have passed it on to our deer managers. State park managers will continue to review their deer hunting programs with the goal of getting our herds where they belong and/or keeping them there. Healthy forests are not just pleasant places to visit, they are critical for the well being of our watersheds, wildlife, plant communities and overall ecology. Once again, thank you for your email. Governor O’Malley appreciates hearing from you, and on his behalf, I thank you for your interest in this very important issue. If you need further assistance, please feel free to discuss this matter with Pete Jayne, Associate Director for Game Management for the Wildlife and Heritage Service, at 410-827-8612 ext 104, or email [email protected]. He will be happy to assist you.

Sincerely, John R. Griffin Secretary

cc: Paul Peditto, Director, Wildlife and Heritage Service Pete Jayne, Associate Director, Wildlife and Heritage Service Nita Settina, Superintendent, Maryland Park Service

IQ No: 315217/CorrNo: 12777-2011

Marilandica Spring 2012 page 4 MNPS Field Trip Plant Lists: A Continuing Project

In the spring of 2007, my husband and I planned a vacation in Wash- USDA nomenclature. ington State to visit family and—of course—to do some hiking and Now we have a very smooth operation. A plant list is sent to me by botanizing. We had previously taken wonderful trips to Mt. Rainier the leader or the recorder soon after a field trip. I check it over and send and Olympic National Park, so this time we planned to take the ferry it on to Bob, who puts it through his magical program, adding family to the San Juan Islands. On a whim, I visited the Washington Native names and footnotes with synonyms. Bob sends the list back to me to Plant Society web page and, to my delight, I found dozens of plant lists check for accuracy. I evaluate the likelihood of each species at the from their field trips. I printed several lists from San Juan County, and particular location and sometimes ask questions of the leader and/or I was on my way. When we arrived, I pulled out two lists for nearby the recorder. After several more rounds of changes, new footnoting, sites, and easily found many of the species listed! and other picky things, we post the list on the MNPS web site. On our return home, I had a plan to post lists from MNPS field Some lists cannot be posted, such as those from private property or trips on our web site. I thought such lists would be useful to field trip very sensitive sites. We have obtained permission to post lists from leaders and participants. I also hoped they might become valuable certain Nature Conservancy preserves and the Izaak Walton League records for future botanists. Well, my project ran into stumbling blocks property in Montgomery County. right away. e main problem was to establish a common format for Below is a combined list from five field trips to the Hereford area the lists. People were sending many different kinds of lists: some with of Gunpowder Falls State Park. is area is particularly noted for common names only, some with no information on who was the leader species such as Hemlock, Ninebark, and Wood Betony that are and who the recorder. Still, the project got started and we had posted typically found farther north or at higher elevations. ten lists by the end of 2008. You can download this list and other lists from our web site. By spring 2009, it was clear there were more hurdles: 1) which Currently, there are 45 lists posted. Go to mdflora.org, and click on authority to use for scientific names; and 2) my appalling lack of “Plant Lists.” Please consider volunteering to be a recorder for MNPS computer skills. So we established a “field trip list committee” to restart field trips. All you need to do is stay near the leader to get plant identi- the project: Ginny and Bob Yacovissi, Heidi Pringle, Kirsten Johnson fications and then check your list with the leader after the trip (don’t and myself. Meeting several times during the summer, the committee worry about the exact spelling). It’s a great way to learn the plants and decided to adopt the USDA database terminology since it is an easily help MNPS keep records at the same time. Leaders, please ask for available and recognized authority. And my lack of computer skills recorders on your field trips! suddenly didn’t matter when Bob offered his expertise as well as access ~ Cris Fleming, former MNPS President, current board member, to a program that picks up spelling mistakes and discrepancies with and author of Finding Wildflowers in the Washington-Baltimore Area

Gunpowder Falls State Park (Hereford Area) Baltimore County, MD We offer these lists to individuals and groups to enhance the enjoyment and study of plants of different locations in Maryland and nearby states. eir accuracy has not been verified by the Maryland Native Plant Society.

Acer negundo Boxelder Aceraceae Adiantum pedatum Acer rubrum Red maple Aceraceae Actaea racemosa 1 Black cohosh Ranunculaceae Adiantum pedatum Maidenhair fern Pteridaceae gryposepala Agrimony Agrimonia parviflora Small-flowered agrimony Rosaceae Agrimonia striata Woodland agrimony Rosaceae Alliaria petiolata 2 Garlic mustard Brassicaceae Ambrosia trifida Giant ragweed Amelanchier sp. Shadbush Rosaceae Amphicarpaea bracteata Hog peanut Fabaceae Asarum canadense Anemone quinquefolia Wood anemone Ranunculaceae Antennaria plantaginifolia Plantain-leaved pussytoes Asteraceae Apocynum cannabinum Indian hemp Apocynaceae Aquilegia canadensis Wild columbine Ranunculaceae Aralia nudicaulis Wild sarsaparilla Araliaceae Aralia racemosa Spikenard Araliaceae Arisaema triphyllum Jack-in-the-pulpit Araceae Asarum canadense Wild ginger Aristolochiaceae Asplenium platyneuron Ebony spleenwort Aspleniaceae Asplenium rhizophyllum 3 Walking fern Aspleniaceae

Marilandica Spring 2012 Maryland Native Plant Society Plant List page 5 Athyrium felix-femina Lady fern Dryopteridaceae Asplenium rhizophyllum Berberis thunbergii Japanese barberry Berberidaceae Betula lenta Black birch Betulaceae cylindrica False nettle Botrychium virginianum Rattlesnake fern Ophioglossaceae Cardamine bulbosa Spring cress Brassicaceae Cardamine concatenata4 Cut-leaved toothwort Brassicaceae Cardamine hirsuta Hairy bittercress Brassicaceae Cardamine pensylvanica bittercress Brassicaceae Carex pensylvanica Pennsylvania sedge Cyperaceae Carpinus caroliniana Hornbeam Betulaceae Carya alba 5 Mockernut hickory Juglandaceae Carya cordiformis Bitternut hickory Juglandaceae Carya glabra Pignut hickory Juglandaceae Castanea dentata Chestnut Fagaceae Celastrus orbiculatus Oriental bittersweet Celastraceae Chelone glabra Turtlehead Scrophulariaceae Chimaphila maculata Spotted wintergreen Pyrolaceae Circaea lutetiana 6 Enchanter's nightshade Onagraceae Claytonia virginica Spring beauty Portulacaceae Carex pensylvanica sp. Tree Collinsonia canadensis Horsebalm Lamiaceae Commelina communis Asiatic dayflower Commelinaceae Cornus alternifolia Alternate-leaved dogwood Cornaceae Cornus florida Flowering dogwood Cornaceae Corydalis flavula Yellow corydalis Fumariaceae Corylus americana Hazelnut Betulaceae Cryptotaenia canadensis Honewort Cunila origanoides Dittany Lamiaceae Cypripedium acaule Pink lady's-slipper Orchidaceae Cystopteris protrusa Fragile fern Dryopteridaceae Cypripedium acaule Dennstaedtia punctilobula Hayscented fern Dennstaedtiaceae Deparia acrostichoides 7 Silvery glade fern Dryopteridaceae Desmodium nudiflorum Naked-flowered tick trefoil Fabaceae Desmodium paniculatum Panicled tick trefoil Fabaceae Dicentra cucullaria Dutchman's breeches Fumariaceae Dichanthelium clandestinum 8 Deer tongue grass Poaceae Dioscorea quaternata Wild yam Dioscoreaceae Dryopteris carthusiana Spinulose wood fern Dryopteridaceae Dryopteris intermedia Intermediate wood fern Dryopteridaceae Dryopteris marginalis Marginal wood fern Dryopteridaceae Echinocystis lobata Burr cucumber Cucurbitaceae Elaeagnus umbellata Autumn olive Elaeagnaceae Dennstaedtia punctiloba Epigaea repens Trailing arbutus Equisetum arvense Common horsetail Equisetaceae Erigeron philadelphicus Common fleabane Asteraceae Erigeron pulchellus Robin's plantain Asteraceae Erythronium americanum Trout lily Liliaceae Euonymus alatus Burning bush Celastraceae Eupatoriadelphus fistulosus 9 Joe-Pye weed Asteraceae Euphorbia corollata Flowering spurge Euphorbiaceae Eurybia divaricata 10 White wood aster Asteraceae Fragaria vesca Wood strawberry Rosaceae Galearis spectabilis 11 Showy orchis Orchidaceae Dicentra cucullaria Galium aparine Cleavers Rubiaceae

Marilandica Spring 2012 Maryland Native Plant Society Plant List page 6 Galium circaezans Wild licorice Rubiaceae Gaultheria procumbens Wintergreen Ericaceae Gaylussacia frondosa Dangleberry Ericaceae Gentiana clausa Closed gentian Gentianaceae Geranium maculatum Wild geranium Geraniaceae Geum canadense White avens Rosaceae Glechoma hederacea Ground ivy Lamiaceae Goodyera pubescens Downy rattlesnake-plantain Orchidaceae Hamamelis virginiana Witchhazel Hamamelidaceae Erigeron philadelphicus Helianthus divaricatus Woodland sunflower Asteraceae Hemerocallis fulva Daylily Liliaceae Hepatica nobilis var. obtusa 12 Round-lobed hepatica Ranunculaceae Heuchera americana Alumroot Saxifragaceae paniculatum Panicled hawkweed Asteraceae Hieracium scabrum Rough hawkweed Asteraceae Hieracium venosum Rattlesnake weed Asteraceae Houstonia caerulea Bluets Rubiaceae Huperzia lucidula 13 Shining clubmoss Hydrangea arborescens 14 Wild hydrangea Hydrangeaceae Hydrophyllum virginianum Virginia waterleaf Hydrophyllaceae Galearis spectabilis Ilex verticillata Winterberry Aquifoliaceae Impatiens capensis Jewelweed Balsaminaceae Isotria verticillata Large whorled pogonia Orchidaceae Juglans cinerea Butternut Juglandaceae Juglans nigra Black walnut Juglandaceae Kalmia latifolia Mountain laurel Ericaceae Laportea canadensis Wood nettle Urticaceae Ligustrum vulgare Common privet Oleaceae Lilium superbum Turk's-cap lily Liliaceae Lindera benzoin Spicebush Lauraceae Liriodendron tulipifera Tuliptree Magnoliaceae Geranium maculatum Lobelia inflate Indian tobacco Campanulaceae Lonicera japonica Japanese honeysuckle Caprifoliaceae Lupinus perennis Wild lupine Fabaceae digitatum 15 Ground pine Lycopodiaceae Lycopodium obscurum 16 Tree clubmoss Lycopodiaceae Lyonia ligustrina Maleberry Ericaceae Maianthemum canadense Canada mayflower Liliaceae Maianthemum racemosum 17 False Solomon's seal Liliaceae Medeola virginiana Indian cucumber-root Liliaceae Menispermum canadense Moonseed Menispermaceae Microstegium vimineum Japanese stiltgrass Poaceae Mitchella repens Partridgeberry Rubiaceae Helianthus divaricatus Mitella diphylla Miterwort Saxifragaceae Nasturtium officinale Watercress Brassicaceae Onoclea sensibilis Sensitive fern Dryopteridaceae Ornithogalum umbellatum Star-of-Bethlehem Liliaceae claytonii Sweet cicely Apiaceae Osmorhiza longistylis Aniseroot Apiaceae Osmunda cinnamomea Cinnamon fern Osmundaceae Osmunda claytoniana Interrupted fern Osmundaceae Ostrya virginiana Hophornbeam Betulaceae Oxalis violacea Violet wood sorrel Oxalidaceae Packera aurea 18 Golden ragwort Asteraceae Hydrophyllum virginianum Panax trifolius Dwarf ginseng Araliaceae

Marilandica Spring 2012 Maryland Native Plant Society Plant List page 7 Parthenocissus quinquefolia Virginia creeper Vitaceae Pedicularis canadensis Wood betony Scrophulariaceae Physocarpus opulifolius Ninebark Rosaceae Pilea pumila Clearweed Urticaceae Platanus occidentalis Sycamore Platanaceae Podophyllum peltatum Mayapple Berberidaceae Polemonium reptans Greek valerian Polemoniaceae Polygonatum biflorum Solomon's seal Liliaceae Polygonum cuspidatum Japanese knotweed Polygonaceae Lindera benzoin Polygonum perfoliatum Mile-a-minute Polygonaceae Polygonum virginianum 19 Jumpseed Polygonaceae Polypodium virginianum Common polypody Polypodiaceae Polystichum acrostichoides Christmas fern Dryopteridaceae Polytrichum sp. Haircap moss Polytrichaceae Potentilla canadensis Dwarf cinquefoil Rosaceae Potentilla simplex Common cinquefoil Rosaceae Prenanthes trifoliolata Tall rattlesnake-root Asteraceae Prunella vulgaris Heal-all Lamiaceae Prunus serotina Black cherry Rosaceae Pycnanthemum muticum Mountain mint Lamiaceae Packera aurea Pycnanthemum virginianum Virginia mountain mint Lamiaceae elliptica Shinleaf Pyrolaceae Quercus alba White oak Fagaceae Quercus prinus Chestnut oak Fagaceae Quercus rubra Northern red oak Fagaceae Ranunculus abortivus Aborted buttercup Ranunculaceae Rhododendron periclymenoides 20 Pinxterflower Ericaceae Rosa multiflora Multiflora rose Rosaceae Rubus allegheniensis Blackberry Rosaceae Rubus phoenicolasius Wineberry Rosaceae Rudbeckia laciniata Tall coneflower Asteraceae Physocarpus opulifolius Salvia lyrata Lyre-leaved sage Lamiaceae Sambucus nigra 21 Elderberry Caprifoliaceae Sanguinaria canadensis Bloodroot Papaveraceae Sanicula odorata 22 Clustered snakeroot Apiaceae Sassafras albidum Sassafras Lauraceae Saxifraga pensylvanica Swamp saxifrage Saxifragaceae Saxifraga virginiensis Early saxifrage Saxifragaceae Silene caroliniana Wild pink Caryophyllaceae Sisyrinchium angustifolium Blue-eyed grass Iridaceae Smilax herbacea Carrion flower Smilacaceae Smilax rotundifolia Greenbrier Smilacaceae Podophyllum peltatum bicolor Silverrod Asteraceae Solidago caesia Blue-stemmed goldenrod Asteraceae Solidago rugosa Rough-stemmed goldenrod Asteraceae Spiraea japonica Japanese spirea Rosaceae Stellaria graminea Common stitchweed Caryophyllaceae Stellaria pubera Star chickweed Caryophyllaceae Symphyotrichum lateriflorum 23 Calico aster Asteraceae Symphyotrichum novae-angliae 24 New England aster Asteraceae Symplocarpus foetidus Skunk cabbage Araceae alictrum dasycarpum Purple meadowrue Ranunculaceae alictrum dioicum Early meadowrue Ranunculaceae alictrum pubescens 25 Tall meadowrue Ranunculaceae Rhododendron periclymenoides alictrum thalictroides 26 Rue anemone Ranunculaceae

Marilandica Spring 2012 Maryland Native Plant Society Plant List page 8 elypteris noveboracensis New York fern elypteridaceae Tiarella cordifolia Foam flower Saxifragaceae Toxicodendron radicans Poison ivy Anacardiaceae Tradescantia virginiana Spiderwort Commelinaceae Trillium cernuum Nodding trillium Liliaceae Tsuga canadensis Hemlock Pinaceae Ulmus rubra Slippery elm Ulmaceae Uvularia perfoliata Perfoliate bellwort Liliaceae Uvularia sessilifolia Sessile-leaved bellwort Liliaceae Sambucus nigra Vaccinium angustifolium Narrow-leaved blueberry Ericaceae Vaccinium corymbosum Highbush blueberry Ericaceae Vaccinium fuscatum 27 Black highbush blueberry Ericaceae Vaccinium pallidum 28 Lowbush blueberry Ericaceae Vaccinium stamineum Deerberry Ericaceae Veratrum viride False hellebore Liliaceae Veronica arvensis Corn speedwell Scrophulariaceae Veronica officinalis Common speedwell Scrophulariaceae Viburnum acerifolium Maple-leaved viburnum Caprifoliaceae Viburnum dentatum Arrowwood Caprifoliaceae Viburnum prunifolium Black haw Caprifoliaceae Sanguinaria canadensis Vinca minor Periwinkle Apocynaceae Viola blanda Sweet white violet Violaceae Viola pubescens 29 Smooth yellow violet Violaceae Viola sororia 30 Common blue violet Violaceae Viola striata Pale violet Violaceae Woodsia obtusa Blunt-lobed cliff fern Dryopteridaceae

1 Cimicifuga racemosa 11 Orchis spectabilis 21 Sambucus canadensis 2 Alliaria officinalis 12 Hepatica americana 22 Sanicula gregaria 3 Camptosorus rhizophyllus 13 Lycopodium lucidulum 23 Aster lateriflorus 4 14 24 Silene caroliniana Dentaria laciniata Hydrangea americana Aster novae-angliae 5 Carya tomentosa 15 Diphasiastrum digitatum 25 alictrum polygamum 6 Circaea quadrisulcata 16 Dendrolycopodium obscurum 26 Anemonella thalictroides 7 Athyrium thelypterioides 17 Smilacina racemosa 27 Vaccinium atrococcum 8 Panicum clandestinum 18 Senecio aureus 28 Vaccinium vacillans 9 Eupatorium fistulosum 19 Tovara virginiana 29 Viola pensylvanica 10 Aster divaricatus 20 Rhododendron nudiflorum 30 Viola papilionacea

alictrum thalictroides is is a combined record of plant lists developed during five different field trips to Gunpow- der Falls State Park (Hereford Area) in 2006, 2007, 2008 and 2009. Field trips were led by Dwight Johnson. Plant lists by Dwight Johnson, Kirsten Johnson, Jane Hill, Heidi Pringle, and Ellen Scarfutti. Nomenclature follows the USDA Plant Database at http://plants.usda. gov (January 2011). Synonyms are footnoted for some species.

Photos by Kirsten Johnson except Asplenium rhizophyllum, Adiantum pedatum and Dennstaedtia punctiloba by Viola blanda Carole Bergmann.

Marilandica Spring 2012 Maryland Native Plant Society Plant List page 9 is is the second of a series of three peer-reviewed articles on floral visitors and phenology of three rare Potomac Gorge plant species. In supporting this project, the National Park Service hoped to learn whether limitation contributes to the rarity of these plants.

Fruit Production and Phenology of covillei S. Watson (Hydrophyllaceae) in the Potomac Gorge Area of Maryland and Virginia

Edward M. Barrows1, Aaron F. Howard2, and Brent W. Steury3

1Laboratory of Entomology and Biodiversity, Department of Biology and the Georgetown University Center for the Environment, Box 571229, Georgetown University, Washington, D.C., 20057-1229

2Laboratory of Entomology and Biodiversity, Department of Biology, and the Georgetown University Box 571229, Georgetown University, Washington, D.C., 20057-1229

3National Park Service, George Washington Memorial Parkway, Turkey Run Park Headquarters, McLean, VA 22101

Abstract lacea (Nutt.) Constance, which they list as endangered in each state (Sewell, 2003; Sewell and Vincent, 2006). Phacelia covillei also occurs e goals of this study are to provide information on selected aspects in the District of Columbia, , and , where officials of the phenology of Phacelia covillei S. Watson in the Potomac Gorge have not assigned it a conservation status. us far, researchers have of the U.S. Mid-Atlantic Region, to ascertain whether this species primarily studied this plant’s morphology, , geographical requires pollination by animals in fruit production in Turkey Run distribution, and response to day length and temperature; however, Park, Virginia, and to examine this species’ flower, fruit, and they have undertaken only limited research on its phenology and production in relation to plant size. Observations and experiments almost no research on its reproductive biology. is species (n = 14 indicate that in our study area, P. covillei sprout in September chromosomes) is morphologically similar to P. ranunculacea (n = 6; and October; plants overwinter as rosettes; and flower, fruit, and Sewell, 2003). senesce in April through early June. Study plants evidently produced no nectar, and insects infrequently visited their flowers. Phacelia Phacelia covillei plants are frequently winter annuals and infrequently covillei sets seeds through intrafloral autogamy, but can also set seeds biennials that grow in rich woods and alluvium (Allard, 1940; through interfloral autogamy and xenogamy based on our hand- Fernald, 1950). Phacelia covillei seeds sprout in the fall and plants pollination and pollinator-exclusion experiments. e number of overwinter as rosettes. Many plants grow in areas with a layer of fallen branches and flowers, fruit, and seeds produced per original flower woody-plant leaves and branches. eir stems below their cotyledons per plant varied significantly among plants in three study sites. are often elongated and their cotyledons and basal, regular leaves Flower, fruit, and seed number were positively correlated with branch usually have long petioles, which enable them to rise above the forest number. Proximal flowers on inflorescences produced more seeds per litter into the light (pers. obs.). In late winter and spring, plants original flower than distal ones. develop 1–10 erect and spreading branches up to 3 dm long, and they flower in April and May. Terminal scorpioid , which become Introduction recurved, each bear 1–6 flowers. e small, light blue-violet flowers are 4–5 mm long, usually five-lobed, and tubular-campanulate Phacelia is a New World genus in the Hydrophyllaceae with over 130 through almost funnelform. e flowers appear to be adapted for annual, biennial, and perennial species, predominately in western insect, not wind, pollination because their corollas are colorful, and North America (Lawrence, 1951; Stevens, 1961; Jaeger, 1969). Pollen mature anthers and stigmas are inside their corollas rather than far and nectar from many species of Phacelia are important resources outside the perianths as occurs in wind-pollinated flowers such as used by pollinating insects and other animals in many habitats some alictrum spp. including agroecosystems and gardens (Williams and Christian, 1991; Williams, 1997; Farkas and Zajácz, 2007). Researchers have Our goals are to describe selected aspects of P. covillei’s phenology, not studied most of the species in detail to obtain a full understanding determine if this species requires insect pollination in setting fruit; of the phenology and reproductive biology of this environmentally and to examine this species’ flower, fruit, and seed production in and economically important genus. relation to plant size.

e herbaceous plant Phacelia covillei S. Watson (Buttercup Scorpi- Materials and Methods onweed, Coville's Phacelia) is endemic to the and occurs in Maryland and (where authorities list it as We made observations on P. covillei from 2003 through 2009 in the S1, endangered). In , , Virginia, and West Virginia, State Potomac River Gorge in Maryland, Virginia, and the District of Heritage Programs have not distinguished this taxon from P. ranuncu- Columbia (Cohn, 2004; Evans, 2008). Our study plants were in

Marilandica Spring 2012 page 10 seven sites in Montgomery County, Maryland and two sites in Fairfax Floral visitors and fruit set County, Virginia, all within 2.4 km of Glen Echo, Maryland. We investigated plants in an experimental garden (Site ME) of the To ascertain whether P. covillei produces seeds without pollination, we Wehawken Nature Preserve in Bethesda, Maryland; three island sites worked at Sites ME and VM-1. At Site ME, we obtained data from in the Potomac River (Sites MI-1, MI-2, and MI-3); three mainland 11 plants. Because it is difficult to emasculate the tiny flowers at sites (MM-1, MM-2, and MM-3) in Maryland; and two mainland near-ground level, we carefully transplanted the plants into pots. e sites (VM-1 and VM-2) in Virginia. Site MI-1 is a low, level island, timing of bud and flower development of the plants allowed us to and MI-2 and MI-3 are hilly islands. Site MM-1 is on a steep, emasculate one test flower on each of the plants. We also chose a wooded slope near the Potomac River; MM-2 is on one of its rolling non-emasculated flower on each plant as a control flower. We marked banks; MM-3 is along one of its tributaries; VM-1 is on a woodland the two kinds of flowers by tying color-coded thread around their knoll; and VM-2 is on a low, flat floodplain. Study sites had from pedicels. We maintained the plants in a greenhouse free of pollina- hundreds through thousands of P. covillei plants that grew both as tors. In Site V-M1, we bagged nine individual flower buds with small isolated plants and in patches of up to thousands of often closely gauze, pollinator-exclusion bags with 0.5-mm mesh, unbagged them spaced plants. Further, all sites had many non-native, invasive plants, when flowers opened and removed their anthers before they dehisced, including Alliaria petiolata (M. Bieb.) Cavara & Grande, Euonymus rebagged the flowers, and subsequently examined their ovaries to see fortunei (Turcz.) Hand.-Maz., and Lonicera japonica unberg. ese if they developed into fruit. plants are especially threatening to P. covillei in Sites MM-1 and MM-3. To determine if P. covillei sets fruit through intrafloral autogamy, we In mid-April 2007, we transplanted nine plants that were not yet in used 16 individually bagged flowers in Site VM-2, and we used test bloom from Site MM-2 into four plastic pots, 16 cm wide x 16 cm cages (pollinator-exclusion cages) and control cages in Sites VM-1 long x 16 cm deep, and transplanted one plant into the ground in a and VM-2. We enclosed entire plants in the cages. We ran the experi- rocky, partly sunny area in Site ME. In our investigation of P. covillei ment with 10 pairs of plants (each with one control and one test reproduction, we tried to choose individual flowers and plants for plant) at Site VM-1 in 2007 and Site VM-2 in 2008. e paired pollination experiments without bias, except for flowers in Site ME, plants were less than 0.3 m apart. We placed cages over plants before from which we chose a flower from each of the 11 plants that grew in they started flowering in mid-April. Each cage was 20 cm in diameter the garden in 2009. We used SPSS Statistics Version 17 to perform by 35 cm high. A control cage had sides of hexagonal chicken wire statistical tests. with each hexagon about 2.5 wide x 3.5 cm long, and a screen top of 1-mm-mesh, gray, plastic window screen. A test cage had sides of Phenology chicken wire covered with gray, plastic window screen and a screen top. We kept a cage in place by driving three 60-cm-long, bamboo To learn about P. covillei plant development, we marked, observed, stakes into the ground around the cage, and tying twine to the stakes and photographed 10 plants at Site VM-1 in 2007. To determine and against the top of each cage so that the twine held down its top when seeds of P. covillei sprout, we observed plants at Sites ME, MI-2, and kept the cage tightly secured to the ground to stop flying pollina- and MI-3. To determine the leaf number of winter rosettes, we tors from entering test cages, wind from blowing cages down, and observed plants in ME in early January 2008 and 40 plants in each of forest vertebrates from knocking over cages. e test cages did not Sites MM-2, VM-1, and VM-2 in late December 2008. At Site ME, exclude possible pollinating ants and tiny thrips; however, we did not we recorded the number of cotyledons and the number of regular see these insects on or in flowers at any site. To record potential and leaves on each of the 15 plants in three pots on an outdoor nursery actual of P. covillei, we directly observed its flowers for a table that was 1 m above the ground and on eight plants in the total of 20 hr from 8 a.m. through 8:30 p.m. during our study period. ground that had sprouted from seeds produced in 2007. In Sites MM-2, VM-1, and VM-2, we laid out 1.5-m-long, twine transects Flower, fruit and seed production, and plant size and counted the leaves on 40 plants along each transect. To ascertain when plants flower and when anthers dehisce, we made direct obser- To determine the number of seeds that each plant produced, we vations on plants. To learn when stigmas are receptive, we used a collected senesced plants and examined their capsules. We found that hydrogen-peroxide test (Kearns and Inouye, 1993) on 20 flowers. We by examining the morphology of a dried that has dropped its observed 10 flowers before their anthers dehisced and 10 plants after seeds, we could determine the number of seeds it produced. We used their anthers dehisced. Each flower was on a different plant. is test a 2-tailed, paired t-test to test the null hypothesis that there is no assumes that peroxidase activity of a stigma indicates that it is recep- significant difference in fruit number or seed number between tive to pollen. e test involves placing a flower’s stigma into a control and test plants. To address the question, “Does P. covillei solution of 3% hydrogen peroxide in water held within a capillary produce seeds through interfloral autogamy?” we bagged nine flower tube. If oxygen bubbles arise from a stigma, it has peroxidase activity. buds with small gauze bags, unbagged them and removed their To find out when plants set fruit, we made observations at study sites, anthers before they dehisced; then pollinated each flower with pollen and we defined fruit set as swelling of a flower’s ovary. We scored a from a flower from the same plant, rebagged flowers; and subse- plant as having mature seeds when it had one or more open capsules, quently examined their ovaries to see if they developed into fruit. To and we scored a plant as senesced when it was limp, brownish, and investigate whether P. covillei can produce seeds through xenogamy, mostly or fully dry, in May. we bagged 13 flower buds with small gauze bags, unbagged them and removed their anthers before they dehisced, pollinated each flower with pollen from a flower from a different plant, rebagged flowers; and subsequently examined the ovaries to see if they developed into fruit.

Marilandica Spring 2012 page 11 Results and Discussion On 22 May 2007 in Site VM-1, 28 study plants had yellowish, mature-looking capsules, and two plants each had dehisced capsules. Phenology At Site MM-2, capsules released seeds 30.8 ± 0.26 (27–33, N = 23) days after their first flowers opened. In 2007 at Site ME, the soil in the ground was very dry during most of July through mid-October, and eight P. covillei plants sprouted At Site VM-1, 30 plants developed from rosettes without flower buds from seeds in the ground on about 31 October after 2 days of rain. through senesced plants that produced seeds from 14 April through Fifteen seeds from plants transplanted from Site MM-1 into pots in early June while other plants developed around them, and new April 2007 sprouted in continually moist soil in pots on 4 October material accumulated and changed positions in the forest litter 2007. Seeds started sprouting in outdoor pots, in which P. covillei (Figure 1). e plants quickly senesced in mid-May through early self-seeded, on 30 September 2009. On 20 October 2007, we found June during a hot, dry period. Allard (1940) remarked that this no seedlings at Sites MI-2 and MI-3 perhaps because the soil was still species is intolerant of temperatures greater than 21°C and quickly too dry that autumn for seed sprouting. In contrast, Allard (1940), senesces in warm May weather. who studied this species in the Potomac River Gorge, reported that P. covillei seeds sprout in December. Seed longevity of P. covillei remains Floral visitors and fruit set unstudied. Seeds of the related winter annual, P. purshii Buckl., germinate in their first through fourth late summers or autumns and During 20 hr of observation that involved all of our sites, we saw only perhaps later (Baskin and Baskin 1976). Seeds of P. ranunculacea a few insects probe P. covillei flowers. ey were two female Andrena sprout in their first, second, or third autumns, making this species an bees (Andrenidae), one female Lasioglossum bee (Halicitidae), two obligate winter annual (Baskin et al., 1993). Bombylius major L. (Bombyliidae), and one geminatus (Say) and one obscurus Say (Syrphidae). ese are taxa that are In the third week of December 2008, P. covillei plants in Site MM-3 common flower visitors and presumed pollinators of many forest and had a mean of 1.60 ± 0.50 1 SD (range 1–2, 40); VM1, 1.40 ± 0.50 other plants. ese insects flew to 1–5 flowers and then left our (1–2, 40); and VM-2, 2.10 ± 0.90 (1–5, 40) true leaves. VM-2 had observed plants. ey appeared to be seeking nectar from the flowers, significantly more such leaves than both MM-3 and VM-1, which but were not obtaining any. We saw no nectar droplets in 10 flowers did not have significantly different numbers of leaves from one from different plants examined with a 20-X hand lens from 9 a.m. – another (homoscedastic data, P = 0.071, Levene Test; P > 0.05, 1-way 12 p.m. at Site MM-2. We did not see any insects collecting or eating ANOVA, Scheffé Test). pollen from the flowers. ese observations suggest that the flower visitors we saw were ones that were inexperienced with P. covillei Times of early P. covillei flowers in the Potomac River Gorge are 12 flowers, and after they learned that the flowers had no nectar, they no April 2003, 17 April 2004, and 17 April 2005 for Site MI-3; 19 April longer visited such flowers. On 25 April 2009, we watched 32 flowers 2005 and 16 April 2006, MM-2; 23 April 2007, MM-3; and 27 April on 17 plants at Site ME after night fall (8–10 p.m) when the tempera- 2007, VM-1; 22 April 2009, MI-3; and 23 April 2009, ME. Allard ture was 26°C, and the air was still. We saw no nocturnal (1940), who also studied specimens of this species from the Potomac visitors at the flowers. Microscopic examination of 10 of these flowers River Gorge, reported that flowers of this species usually appear after at up to 30-X showed that their anthers had dehisced and all stigmas 20 April and continue through mid-May. is species may be had pollen on them. Nine of the flowers had two stigmas, and one currently blooming earlier in the Gorge due to earlier warm weather flower had three stigmas. We saw no nectar droplets in the flowers. related to global climate change. Flower, fruit and seed production, and plant size By spring 2008, 17 plants were present in Site ME, and they all descended from a single plant planted in that site in 2007. One of the Phacelia covillei did not produce seeds by apomixis. At Site VA-2, plants had one tripetalous, four quadripetalous, and five pentapetal- none of the 10 emasculated flowers produced fruit, and at Site ME, ous flowers; eight plants had quadripetalous and pentapetalous none of the 11 emasculated flowers produced fruit. At Site VA-2, we flowers; and eight plants had only pentapetalous flowers on 28 April found that flowers opened with nondehisced anthers, but at Site ME, 2007. In sum, the plants had one tripetalous, eight quadripetalous, individual flowers opened with either dehisced or nondehisced and 36 pentapetalous flowers. e flowers were sympetalous, and we anthers, causing us to open buds and remove anthers before they saw no evidence that corolla lobes had fallen from corollas. opened to study possible fruit set without pollination.

Anthers dehisce within flower buds up to 2 hr after flowers open, and ree lines of evidence indicate that P. covillei flowers set seeds by anthers have pollen in open flowers when they are 1.09 ± 0.06 (1–2, intrafloral autogamy. First, at Site VA-2, after we removed nonde- N = 23) days old. Within buds, nondehisced anthers range from light hisced anthers from each of 15 flowers, opened its anthers, and yellow through orange. In open flowers, nondehisced anthers are pollinated it with its own pollen, nine of the flowers produced seeds. orange, and dehisced ones are light brown when macroscopically Second, at Site ME, all 11 control flowers produced seeds. ird, the examined. Corollas wilt 2.5 ± 0.11 (2–3, N = 22) days after they open. 20 test plants in pollinator-exclusion cages produced seeds.

All the stigmas from 10 open flowers with closed anthers and 10 open Phacelia covillei also produces seeds via interfloral autogamy. At Site flowers with dehisced anthers produced oxygen bubbles in peroxide VM-2, flowers on three of nine plants that we cross-pollinated with indicating that all 20 stigmas were receptive. Ovaries were initially pollen from flowers of the same plant set seeds. Further, this species markedly expanded 4.9 ± 0.29 (3–7, N = 21) days after anthesis. also produces seeds via xenogamy. At Site VM-2, flowers of five of 15

Marilandica Spring 2012 page 12 plants that we cross-pollinated with pollen from a different conspe- In conclusion, we found that in the Potomac River Gorge, the cific plant produced seeds. Like P. covillei, a related species, Phacelia scientifically and esthetically alluring plant P. covillei exists as seeds purshii Buckl. (Gillett, 1964; Baskin and Baskin, 1976), and possibly from June through early October, its seeds sprout in October, rosettes other Phacelia spp. are self-pollinated, self-fertile winter annuals. overwinter, plants flower in April and May, and fruit are mature and plants senesce in May. Phacelia covillei plants have infrequent flower In 2007 at Site VM-1, 10 control plants had a mean of 4.6 ± 2.9 1 SD visitors, and appear to produce fruit entirely, or almost entirely, via (range 2–11) fruit, and 10 test plants had 5.3 ± 1.6 (3–7) fruit, show- self-pollination without animal pollinators as their pollen vectors. ing no difference in fruit number between groups (t = –0.535; P = Plants in pollinator-exclusion cages, can produce up to 19 seeds per 0.606, 2-tailed, paired t-test). Control plants had 7.9 ± 4.9 (4–19) plant. erefore, at least within the Gorge, P. covillei plants can seeds, and test plants had 5.8 ± 4.4 (1–16) seeds, showing no differ- produce many seeds without animal pollination. Many lines of ence in seed number between groups (t = 1.058; P = 0.318). Control research remain to be undertaken on P. covillei including how the plants had 1.8 ± 0.3 (1–2) seeds, and test plants had 1.6 ± 0.3 (1–2) reproductive biology of other populations may vary from that of the seeds per fruit, showing no difference in seed number per fruit Gorge population. For example, are there populations with nectarif- between groups (t = 1.551; P = 0.155). In 2007 at Site VM-1, both erous flowers that are frequently visited by insect pollinators? How control and test plants had only 1–2 seeds per fruit, but at this does the phenology and reproductive biology of P. covillei differ from location in 2008, we found plants with 1–4-seeded capsules. In 2008 that of the similar-looking P. ranunculacea? at Site VM-2, 10 control plants had a mean of 7.4 ± 3.7 (2–12) fruit, and test plants had 5.3 ± 1.6 (3–7) fruit, showing no significant Acknowledgements difference in fruit number between groups (t = 2.003; P = 0.076, 2-tailed, paired t-test). Control plants had a mean of 17.7 ± 9.1 Georgetown University, the National Park Service (NPS), and the (4–33) seeds, and test plants had 10.4 ± 3.5 (5–15) seeds showing Washington Biologists’ Field Club provided financial support for this significantly more seeds in controls (t = 2.916; P = 0.017). Control study. e NPS provided permits to work in our study sites. Patrick plants had 2.4 ± 0.4 (1.8–2.8) fruit, and test plants had 2.0 ± 0.3 M. Differding and anonymous peer reviewers provided constructive (1.6–2.5) seeds per fruit showing more seeds per fruit in controls (t = criticisms regarding this paper. 5.369; P = 0.016). At both sites, control plants had more seeds and seeds per fruit than test plants. Greater shading of test plants Literature Cited compared to control plants by the full screening of test cages may have reduced test-plant seed set. Allard, H. A. 1940. (Nutt.) Constance, its length of day, temperature reactions, and seasonal adaptations. Casta- e 120 plants from all three study sites combined had 3.3 ± 1.8 nea 5: 94–97. (1–10) branches, 9.4 ± 6.7 (1–37) flowers, 4.9 ± 3.8 (0–19) fruit, 12.8 ± 9.6 (0–57) seeds, 2.7 ± 0.7 seeds per fruit, and 0.3 ± 0.3 seeds Baskin, J. M., and C. C. Baskin. 1976. Some aspects of the autecol- per original flower. Site VM-2 plants had significantly more branches, ogy and population biology of Phacelia purshii. American Midland flowers, fruit, seeds, and seeds produced per flower than plants at the Naturalist 96: 431–442. other two sites (Table 1). Further, our results suggest that plants of Sites VM-1 and VM-2 had more seeds per fruit than plants of Site Baskin, C. C., J. M. Baskin, and E. W. Chester. 1993. Seed germina- MM-2. Flower, fruit, and seed number are positively correlated with tion ecology of two mesic woodland winter annuals, Nemophila branch number for all sites (Table 2, Pearson’s and Spearman’s Rho aphylla and Phacelia ranunculacea (Hydrophyllaceae). Bulletin of the Correlation Tests, P ≤ 0.01, in all 18 correlations). erefore, more Torrey Botanical Club 120: 29–37. branched plants have higher Darwinian fitness (measured as seed number) than less branched ones. Researchers evidently have not Cohn, J. P. 2004. e wildest urban river: Potomac River Gorge. studied the possible effects of space and nutrient availability on BioScience 54: 8–14. branching in this species; however, Allard (1940) found that P. covillei rosettes under lower-light conditions branched more than plants Evans, A. V. 2008. e 2006 Potomac Gorge Bioblitz. Overview and under higher-light conditions in a greenhouse. results of a 30-hour rapid biological survey. Banisteria Number 32: 3–80. At all three sites, the more proximal flowers of an inflorescence, which tend to bloom earlier than the more distal ones, produced more seeds Farkas, Á., and E. Zajácz. 2007. Nectar production for the Hungarian than more distal flowers (Table 3). e possible causes for this honey industry. e European Journal of Plant Science and Biotech- phenomenon in P. covillei are not yet studied, but may include nology 1: 125–151. decreasing nutrient and light availability for flower maturation and seed production as the flowering season progresses and forest canopies Fernald, M. L. 1950. Gray's Manual of Botany, 8th ed. American become denser. Book Company, New York, New York. 1632 pp.

Hydrophyllaceae evidently has a highly conserved breeding system Gillett, G. W. 1964. Genetic barriers in the Cosmanthus phacelia with protandry, maximal stigma receptivity near day 3, and self-com- (Hydrophyllaceae). Rhodora 66: 359–368. patibility (Levy, 1988). Phacelia covillei is an exception to this trend in that its anthers may dehisce within buds, its flowers tend to last only Jaeger, E. C. 1969. Desert Wild Flowers. Stanford University Press, 2 days, and its stigmas are evidently receptive during both days. Stanford, California. 321 pp.

Marilandica Spring 2012 page 13 Kearns, C. A. and D. W. Inouye. 1993. Techniques for Pollination Edward “Edd” Barrows is Professor of Biology and Director of the Biologists. University Press of Colorado, Niwot, Colorado. 583 pp. Laboratory of Entomology and Biodiversity, Environmental Studies, and the Center for the Environment at Georgetown University. Edd’s research Lawrence, G. H. M. 1951. Taxonomy of Vascular Plants. Macmillan focuses on behavior, ecology, evolution, and conservation, usually involv- Company, New York, New York. 823 pp. ing and plants as study organisms. He is also an expert in scientific communication and the author of Animal Behavior Desk Levy, F. 1988. Effect of pollen source and time of pollination on seed Reference: A Dictionary of Animal Behavior, Ecology, and Evolu- production and seed weight in Phacelia dubia and P. maculata tion. (Hydrophyllaceae). American Midland Naturalist 119: 193–198. Aaron F. Howard, Ph.D., is a recent graduate of Georgetown University’s Sewell, M. 2003. Biosystematics of the Phacelia ranunculacea com- Laboratory of Entomology and Biodiversity. He is currently on the faculty plex (Hydrophyllaceae). M.S. thesis, Miami University, Ohio. 60 pp. at two colleges in the Chicago area, where he teaches biology and environ- http://www.ohiolink.edu/etd/view.cgi?miami1071261639 mental science. He is completing papers on plant clonality and the ecology and evolution of floral-display size in two North American milkweed Sewell, M., and M. A. Vincent. 2006. Biosystematics of the Phacelia species. ranunculacea complex (Hydrophyllaceae). Castanea 71: 192–209. Brent Steury is the Natural Resources Program Manager for the national Stevens, W. C. 1961. Kansas Wild Flowers, 2nd ed. University of park known as the George Washington Memorial Parkway, where he Kansas Press, Lawrence, Kansas. 461 pp. currently oversees all aspects of the park’s inventory and monitoring program. His research has focused on the distribution of threatened and Williams, C. S. 1997. Nectar secretion rates, standing crops and endangered vascular plants on the Mid-Atlantic Coastal Plain. Brent has flower choice by bees on Phacelia tanacetifolia. Journal of Apicultural authored 20 journal articles, numerous technical reports, and discovered Research 36: 23–32. more than a dozen vascular plants that were new records for the floras of Maryland, Virginia, or the District of Columbia. In addition to his Williams, I. H., and D. G. Christian. 1991. Observations on Phacelia interest in plants, Brent has published papers on reptiles, amphibians, tanacetifolia Bentham (Hydrophyllaceae) as a food plant for honey moths, bees, and land snails. bees and bumble bees. Journal of Apicultural Research 30: 3–12.

Marilandica Spring 2012 page 14 Table 1. e mean (± 1 SD) number of branches, flowers, fruit, seeds, seeds per flower, and seeds per fruit per Phacelia covillei plant (N = 40 plants per site)1.

Site Plant Characteristic Seed number per Branch number Flower number Fruit number Seed number Seed number 2 2 2 flower number 2 per plant per plant per plant per plant per plant3 per fruit 2.7 ± 1.3 6.8 ± 3.9 3.8 ± 2.3 11.1 ± 6.30 0.4 ± 0.3 3.0 ± 0.7 MM-2 (1–6)a (2–20)a (1–11)a (2–33)a (0.1–1.0)a (1.5–6.3)a

2.6 ± 1.5 6.9 ± 5.5 3.2 ± 2.3 8.5 ± 6.67 0.4 ± 0.3 2.4 ± 0.8 VM-1 (1–7)a (1–27)a (0–19)a (0–34)a (0–1)a (0–4)b

4.7 ± 1.6 14.5 ± 7.2 7.4 ± 4.4 18.8 ± 11.65 0.2 ± 0.1 2.5 ± 0.5 VM-2 (1–10)b (4–37)b (2–19)b (5–57)b (0.1–0.5)b (1.5–3.5)b

1 Ranges are in parentheses after means. Within columns, means that do not share a same letter are significantly different from one another at P ≤ 0.05 (ANOVA, Scheffé Test). 2 We used a square-root transformation on these data to meet the homoscedasticity assumption. 3 We used a cube-root transformation on these data to meet the homoscedasticity assumption.

Table 2. Correlations of flower, fruit, and seed number with branch number per Phacelia covillei plant per site. N = 40 plants per site.

1 Site Correlate with Branch Number Flower number Fruit number Seed number

MM-2 0.876, 0.843 0.782, 0.702 0.673, 0.626

VM -1 0.917, 0.946 0.778, 0.879 0.827, 0.856 VM -2 0.911, 0.914 0.871, 0.881 0.814, 0.847

1 e first coefficient in a pair is for the parametric Pearson’s Correlation Test (r), and the second coefficient is for the nonparametric Spearman’s Correlation Test (rho); P ≤ 0.01 for all correlations.

Table 3. e mean number of seeds per original flower site on infructescences per Phacelia covillei plant1.

Site Inflorescence Flower Position

1 2 3 4 5 6 MM-2 2.21 (40) 1.80 (39) 1.40 (33) 0.64 (21) 0 (7) 0 (1) VM-1 2.28 (40) 1.54 (39) 0.41 (30) 0 (18) 0 (2) VM-2 2.31 (40) 1.41 (40) 0.76 (32) 0.14 (32) 0.12 (17) 0 (3)

1 Flower position-1 is the most proximal position; position-2 is the next most proximal position; and so forth. N = 40 Phacelia covillei plants in each of the three sites. e number within parentheses following a mean is the number of plants from which the mean is calculated. Each mean is an average of the mean number of seeds at each flower position for all racemes of each plant. e number of plants varies because although all plants had a flower at position-1, not all plants had flowers at position-2 and at more distal inflorescence positions.

Marilandica Spring 2012 page 15 a b c

d e f

Figure 1. Development of a small plant of Phacelia covillei at Site VM-1 accompanied by changes in growth of other plants and in litter around it, 14 April – 24 May 2007. Photographs differ in scale so that the entire plant is shown in each photograph. Litter accumulated and moved about during the observation period. (a) 14 April, the expanding rosette of the P. covillei plant. A small shoot of Lonicera japonica unberg (Japanese Honeysuckle) is in the left upper corner, and its base is also in figures b–f. A second small shoot of L. japonica is in the lower bottom area and is also visible in figures b–d. A shoot of Claytonia virginica L. (Virginia Spring-beauty) is in the lower left corner, and is also visible in figures b–d before the shoot disappeared. A plant of Veronica hederifolia L. (Ivy-leafed Speedwell) is in the top middle area of the photograph. Seeds of Liriodendron tulipifera L. (Tuliptree) are in all photographs. (b) 21 April, the P. covillei plant with flower buds (arrow). Claytonia virginica (left corner) is in bloom. (c) 25 April, flowers of the P. covillei plant (arrow). A male inflorescence of Fagus grandiflora Ehrh. (American Beech) is near the lower right side of the P. covillei plant. (d) 2 May, small fruit of the P. covillei plant (arrow). (e) 17 May, large fruit of the P. covillei plant (arrow). A shoot of V. hederifolia is in the upper left corner. (f) 24 May, the totally senesced P. covillei plant with dehisced capsules (arrow). Two brownish petals of L. tulipifera are in the right side of the photograph.

Marilandica Spring 2012 page 16 Conservation Watch

Permits Denied for Charles County Connector

In late February, the Army Corps of Engineers joined the Maryland MDE denied the state Nontidal Wetlands and Waterways Permit in Department of the Environment (MDE) in denying a permit for the November. proposed Cross County Connector extension in Charles County because of its likely effect on Mattawoman Creek. e letter of denial In the Fall 2011 Marilandica, Mattawoman Watershed Society states “the project as proposed would have direct and permanent President Jim Long commented that the “Mattawoman story shows adverse impacts on non-tidal wetlands, waterways, fish and wildlife, how efforts to restore the Bay and protect valuable wetlands resemble recreation and water quality.” e Corps also found that practical the story of Sisyphus, with two steps backward for every step alternatives to the project, such as utilizing existing roads, had not forward.” e recent denial of highway permits was unquestionably a been sufficiently explored. Notably, the US Fish and Wildlife Service step forward, but the fragile Mattawoman Watershed–so important had urged denial of the permit, citing President Obama's May 2009 to the health of the Chesapeake Bay–remains seriously threatened by executive order for federal agencies to take a leadership role in development. For more information, please see mattawomanwater- protecting the Chesapeake Bay and its surrounding wildlife habitats. shedsociety.org. Deer Management in Baltimore County - Update As reported in the Spring 2011 Marilandica, a group of Baltimore e amendment approved by the Council fell short of the petitioners’ County citizens petitioned the County Council to permit regulated recommendation, but it permits culling of deer in County parks deer hunting in the County. e petition was signed by 18 organiza- under Deer Cooperator Programs conducted by licensed wildlife tions representing diverse civic, environmental and recreational management companies under the supervision of the Department of interests, but all concerned about the negative effects of deer over- Natural Resources. population on forest health, as well as Lyme disease and highway safety.

Coming Events

MONTHLY MEETINGS Many MNPS members have thought of the monthly meetings in Montgomery County–usually at the Kensington Library, Knowles Avenue, in Kensington–as the regular meetings of the Maryland Native Plant Society. MNPS’s other chapters hold monthly meetings as well; all the meetings known at press time are listed chronologically. Please see www.mdflora.org for details.

April 17 ~ Tuesday, 7:00 PM May 29, Tuesday ~ 7:30 PM, doors open at 6:30 A Native American Species Becomes a New Fruit Crop Deer Management in Montgomery County Western Mountains Chapter. Location: Appalachian Laboratory, Frostburg Montgomery County. Location: Kensington Library Speaker: R. Neal Peterson, Plant Breeder; on the pawpaw, Asimina Speaker: George Timko, Assistant Deer Project Manager, MD DNR. triloba, North America’s largest edible native fruit. June 19, Tuesday ~ 7:00 PM April 18 ~ Wednesday, 7:00 PM Western Mountains Chapter Native Plant Nurseryman Roundtable Location: Appalachian Laboratory, Frostburg Baltimore Chapter. Location: Cylburn Arboretum greenhouse classroom Speaker: Donna Ford-Wentz, Herbarium, West Virginia University Participating Nurseries: Heartwood, Kollar, and American Native Plants. ey will answer questions and bring plants for sale. June 20, Wednesday ~ 7:30 PM Saving Wild Plants at Meadowlark Garden April 24 ~ Tuesday, 7:30 PM, doors open at 6:30 Baltimore Chapter. Location: Druid Hill’s Rawlings Conservatory. The Natural History of Oak-Feeding Insects in Maryland Speaker: Keith Tomlinson, Manager, Meadowlark Botanical Gardens. Montgomery County. Location: Kensington Library Speaker: John Lill, Assoc. Prof. of Biology, George Washington Univ. June 26, Tuesday ~ 7:30 PM, doors open at 6:30 Oaks of the Mid-Atlantic May 16, Wednesday ~ 7:00 PM Montgomery County. Location: Kensington Library Maryland’s Orchids Speaker: Rod Simmons Baltimore Chapter. Location: Irvine Nature Center, Owings Mills Speaker: Carol Allen, Horticulturist and Orchid Enthusiast.

Marilandica Spring 2012 page 17 Coming Events

SPRING FIELD TRIPS ese are the field trips scheduled at press time. For up to date news of MNPS field trips and activities please see our website, www.mdflora.org and find us at meetup.com. Unless otherwise indicated, MNPS field trips are generally geared to adults. Please see the information provided for individual field trips, some of which may welcome children. If you have questions, feel free to contact the field trip leader.

April 21 ~ Saturday, 10:00 AM – 1:00 PM May 6 ~ Sunday, 10:00 AM – 2:00 PM Spring Wildflowers at Governor Bridge Natural Area Civil War Fort Sites in Washington: Fort Chaplin Leader: Karyn Molines Leaders: Mary Pat Rowan and Lou Aronica We’ll walk through a diversity of habitats and identify abundant spring Always a very special trip in May, we see wild flowers which appear only wildflowers. this month. We will also see a variety of native shrubs, which are unusual in a terrace gravel upland forest. April 21 ~ Saturday, Time tba Two events at Druid Hill Park: Druid Hill Oaks and May 6 ~ Sunday, 9:00 AM – 12:00 PM Landscaping with Native Plants to Attract Wildlife Uncultivated: Exploring Wild Urban Plants, location tba Leaders: Chris Partain and Glenda Weber Leaders: Lynn Cazabon and Chris Partain In conjunction with Baltimore’s EcoFest, MNPS will offer a tour of Artist Lynn Cazabon will discuss her ongoing project, Uncultivated, Druid Hill’s majestic oaks. while demonstrating how she chooses sites and photographs the plants.

April 22 ~ Sunday, 10:00 AM – 1:00 PM May 19 ~ Sunday, 10:00 AM – 3:00 PM Leakin Park Gunpowder State Park Leaders: Dwight and Kirsten Johnson Leader: Dwight Johnson Part of Baltimore’s Green Week, this walk explores spring wildflowers of Explore oaks and late spring wildflowers on the Gunpowder in Leakin Park. Hereford. We will explore downstream in the morning, return to lunch at our cars, then go upstream in the afternoon. April 24 ~ Tuesday, 12:00- 12:45 PM Jones Falls at the Inner Harbor May 20 ~ Sunday, 10:00 AM – 1:00 PM Leader: Drew Brown Piney Orchard Nature Preserve A Baltimore’s Green Week event, this is a lunchtime walk along the Leaders: Beth Johnson and Chris Partain Inner Harbor and a discussion of discussing urban stream ecology. Piney Orchard Nature Preserve in Anne Arundel County, the result of wetland restoration efforts, includes wetlands, woodlands, open grassy April 28 ~ Saturday, 10:00 AM- 2:00 PM areas, sandy dry stream beds, and freshwater ponds. We will search for Carderock Recreation Area Woodlands and Tow Path butterflies, dragonflies, damselflies, and plants. Leader: Marney Bruce We will be walking in the woods, on the tow path, and occasionally June 3 ~ Sunday, 10:00 AM – 1:00 PM crossing streams and climbing hills, identifying abundant wildflowers. Oaks of Sugarloaf Mountain Leader: Melanie Choukas-Bradley April 28 ~ Saturday, 10:00 AM – 12:00 PM A leisurely hike to the summit of Sugarloaf Mountain. We will see Herring Run Park several oak species. Mountain laurel and several early summer wildflow- Leader: Vince Vizachero ers will also be in bloom. Online registration is required.

April 29 ~ Sunday, 10:00 AM – 1:00 PM June 17 ~ Sunday, 10:00 AM – 1:00 PM Severn Run Natural Environmental Area Soldiers Delight Natural Environment Area Leader: Chris Partain Leader: Chris Partain is year we will explore the Severn Run Natural Environmental Area Chris Partain will lead the trip as part of the year’s exploration of this through the seasons. is botanically-rich spot, in northern Anne Arundel special area. Serpentine barrens support a unique flora, which is adapted County, is an undeveloped park protecting the Severn Run Watershed. to low essential nutrients but high concentrations of heavy metals. Some scrubby oak species here are blackjack and post oaks. May 1 ~ Tuesday, 6:00 PM – 8:00 PM Near Enchanted Forest Area of Ellicott City June 23 ~ Saturday, 9:00 – 11:00 AM Leader: Heidi Pringle Oaks Overhead, Garrett County is is a small stream valley, and a climb down overgrown riprap rocks Leader: Liz McDowell, MNPS, and Wade Dorsey, Savage River State is necessary to enter the forest. Please see details and register at Forest Manager mdflora.org. Limit of 25 participants. During this leisurely walk Wade will highlight various oak species, and Liz will identify the various plants that inhabit the understory.

Marilandica Spring 2012 page 18 Become a member, join online: www.mdflora.org.

A Publication of the Maryland Native Plant Society

Maryland Native Plant Society PO Box 4877 Silver Spring, MD 20914

Year of the Oak

In this Issue Letter from the President pg 1 Oak in Focus 2 Letter to Governor 3 Photo Essay 3 Letter from Governor 4 MNPS Field Trip Plant Lists 5 Barrows, et al. 10 Conservation Watch 17 Upcoming Meetings 17 Upcoming Field Trips 18

Society Plant www.mdflora.org

j Organization $50 j Lifetime $350 PO Box 4877 www.mdflora.org Nativ e Silver Spring, MD 20914 Silver Spring, Maryland Native Plant Society in Maryland, to which donations are tax deductible. in Maryland, j Individual $25 j Double $35 j Student/Limited Income $15 j Optional Donation $ Enclosed $ Total Annual Membership Dues Make checks payable to: Maryland Native Plant Society or MNPS to: checks payable Make You can also join on-line on our website You Maryland Native Plant Society, Inc. is a Section 501(c)(3) charitable organization incorporated is a Section 501(c)(3) charitable organization incorporated Inc. Maryland Native Plant Society, Member 1 ______Member 2 ______Street ______City ______State ______Co _____ Zip______Phone (S) Day______Eve ______Cell ______Email ______j I do not wish to receive emails